Project 3134: B. M. Gee, J. J. Bevitt, U. Garbe, R. R. Reisz. 2019. New material of the ‘microsaur’ Llistrofus from the cave deposits of Richards Spur, Oklahoma and the paleoecology of the Hapsidopareiidae. PeerJ. 7:e6327.
Abstract
The Hapsidopareiidae is a group of ‘microsaurs’ characterized by a substantial reduction of several elements in the cheek region that results in a prominent, enlarged temporal emargination. The clade comprises two markedly similar taxa from the early Permian of Oklahoma, Hapsidopareion lepton and Llistrofus pricei, which have been suggested to be synonymous by past workers. Llistrofus was previously known solely from the holotype found near Richards Spur, which consists of a dorsoventrally compressed skull in which the internal structures are difficult to characterize. Here we present data from two new specimens of Llistrofus. This includes data collected through the use of neutron tomography, which revealed important new details of the palate and the neurocranium. Important questions within ‘Microsauria’ related to the evolutionary transformations that likely occurred as part of the acquisition of the highly modified recumbirostran morphology for a fossorial ecology justify detailed reexamination of less well-studied taxa, such as Llistrofus. Although this study eliminates all but one of the previous features that differentiated Llistrofus and Hapsidopareion, the new data and re-description identify new features that justify the maintained separation of the two hapsidopareiids. Llistrofus possesses some of the adaptations for a fossorial lifestyle that have been identified in recumbirostrans but with a lesser degree of modification (e.g., reduced neurocranial ossification, mandibular modification). Incorporating the new data for Llistrofus into an existing phylogenetic matrix maintains the Hapsidopareiidae’s (Llistrofus + Hapsidopareion) position as the sister group to Recumbirostra. Given its phylogenetic position, we contextualize Llistrofus within the broader ‘microsaur’ framework. Specifically, we propose that Llistrofus may have been fossorial but was probably incapable of active burrowing in the fashion of recumbirostrans, which had more consolidated and reinforced skulls. Llistrofus may represent an earlier stage in the step-wise acquisition of the derived recumbirostran morphology and paleoecology, furthering our understanding of the evolutionary history of ‘microsaurs.’Read the article »
Article DOI: 10.7717/peerj.6327
Project DOI: 10.7934/P3134, http://dx.doi.org/10.7934/P3134
This project contains | Matrices |
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Download Project SDD File | Total scored cells: 26166 Total media associated with cells: 0 Total labels associated with cell media: 0 |
Characters | |
Total characters: 597 Total characters with associated media: 0 Total characters with media with labels: 0 Total character states: 1008 Total character states with associated media: 0 Total character states with media with labels:0 Total unordered/ordered characters:597/0 |
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MorphoBank Project 3134
MorphoBank Project 3134
- Creation Date:
18 March 2018 - Publication Date:
27 December 2018 - Project views: 43518
- Media downloads: 1726
- Matrix downloads: 4
Authors' Institutions
- University of Toronto
Members
member name | taxa | specimens | media | chars | character
| cell scorings (scored, NPA, "-") | cell
| rules | ||||||
Bryan Gee Project Administrator | 1 | 1 | 3 | 0 | 0 | 0 | 0 (0, 0, 0) | 0 | 0 | 0 | ||||
MorphoBank Curator Full membership | 112 | 0 | 0 | 597 | 0 | 0 | 26166 (24352, 0, 1814) | 0 | 0 | 0 |
Taxonomic Overview for Matrix 'M25827' (58 Taxa)
taxon | unscored cells |
scored cells |
no cell support |
NPA cells |
"-" cells | cell images | labels on cell images |
member access |
[1] † Archeria crassidisca Last Modified in 01/25/19 | 50 | 309 | 308 | 0 | 12 | 0 | 0 | 2 |
[2] † Crassigyrinus scoticus Last Modified in 01/25/19 | 82 | 281 | 281 | 0 | 7 | 0 | 0 | 2 |
[3] † Acanthostega gunnari Last Modified in 01/25/19 | 15 | 350 | 349 | 0 | 6 | 0 | 0 | 2 |
[4] † Eusthenopteron foordi Last Modified in 01/25/19 | 154 | 196 | 196 | 0 | 20 | 0 | 0 | 2 |
[5] † Ichthyostega spp. Last Modified in 01/25/19 | 57 | 307 | 305 | 0 | 8 | 0 | 0 | 2 |
[6] † Panderichthys rhombolepis Last Modified in 01/25/19 | 263 | 107 | 107 | 0 | 0 | 0 | 0 | 2 |
[7] † Pederpes finneyae Last Modified in 01/25/19 | 163 | 203 | 203 | 0 | 4 | 0 | 0 | 2 |
[8] † Tiktaalik rosaea Last Modified in 01/25/19 | 91 | 245 | 244 | 0 | 35 | 0 | 0 | 2 |
[9] † Ventastega curonica Last Modified in 01/25/19 | 125 | 241 | 241 | 0 | 4 | 0 | 0 | 2 |
[10] † Whatcheeria deltae Last Modified in 01/25/19 | 86 | 278 | 277 | 0 | 7 | 0 | 0 | 2 |
[11] † Ymeria denticulata Last Modified in 01/25/19 | 267 | 102 | 101 | 0 | 2 | 0 | 0 | 2 |
[12] † Proterogyrinus scheelei Last Modified in 01/25/19 | 46 | 317 | 313 | 0 | 11 | 0 | 0 | 2 |
[13] † Balanerpeton woodi Last Modified in 01/25/19 | 77 | 286 | 286 | 0 | 7 | 0 | 0 | 2 |
[14] † Dendrerpeton acadianum Last Modified in 01/25/19 | 46 | 318 | 316 | 0 | 8 | 0 | 0 | 2 |
[15] † Pantylus cordatus Last Modified in 01/25/19 | 11 | 340 | 339 | 0 | 20 | 0 | 0 | 2 |
[16] † Cardiocephalus peabodyi Last Modified in 01/25/19 | 206 | 157 | 156 | 0 | 8 | 0 | 0 | 2 |
[17] † Huskerpeton englehorni Last Modified in 01/25/19 | 107 | 243 | 243 | 0 | 20 | 0 | 0 | 2 |
[18] † Pelodosotis elongatum Last Modified in 01/25/19 | 51 | 301 | 300 | 0 | 19 | 0 | 0 | 2 |
[19] † Nannaroter mckinzei Last Modified in 01/25/19 | 165 | 186 | 186 | 0 | 19 | 0 | 0 | 2 |
[20] † Micraroter erythrogeios Last Modified in 01/25/19 | 94 | 258 | 256 | 0 | 20 | 0 | 0 | 2 |
[21] † Eocaecilia micropodia Last Modified in 01/25/19 | 52 | 262 | 261 | 0 | 57 | 0 | 0 | 2 |
[22] † Carrolla craddocki Last Modified in 01/25/19 | 119 | 211 | 211 | 0 | 40 | 0 | 0 | 2 |
[23] † Quasicaecilia texana Last Modified in 01/25/19 | 211 | 141 | 141 | 0 | 18 | 0 | 0 | 2 |
[24] † Llistrofus pricei Last Modified in 01/25/19 | 91 | 254 | 254 | 0 | 25 | 0 | 0 | 2 |
[25] † Aletrimyti gaskillae Last Modified in 01/25/19 | 86 | 255 | 255 | 0 | 29 | 0 | 0 | 2 |
[26] † Dvellecanus carrolli Last Modified in 01/25/19 | 107 | 237 | 237 | 0 | 26 | 0 | 0 | 2 |
[27] † Rhynchonkos stovalli Last Modified in 01/25/19 | 105 | 239 | 239 | 0 | 26 | 0 | 0 | 2 |
[28] † Captorhinus laticeps Last Modified in 01/25/19 | 15 | 317 | 316 | 0 | 39 | 0 | 0 | 2 |
[29] † Brachydectes newberryi Last Modified in 01/25/19 | 14 | 327 | 326 | 0 | 30 | 0 | 0 | 2 |
[30] † Greererpeton burkemorani Last Modified in 01/25/19 | 15 | 338 | 337 | 0 | 18 | 0 | 0 | 2 |
[31] † Seymouria baylorensis Last Modified in 01/25/19 | 16 | 341 | 338 | 0 | 16 | 0 | 0 | 2 |
[32] † Limnoscelis paludis Last Modified in 01/25/19 | 17 | 335 | 334 | 0 | 19 | 0 | 0 | 2 |
[33] † Branchiosauridae Last Modified in 01/25/19 | 208 | 156 | 154 | 0 | 8 | 0 | 0 | 2 |
[34] † Micromelerpetontidae Last Modified in 01/25/19 | 212 | 152 | 151 | 0 | 7 | 0 | 0 | 2 |
[35] † Tersomius texensis Last Modified in 01/25/19 | 158 | 212 | 206 | 0 | 6 | 0 | 0 | 2 |
[36] † Ecolsonia cutlerensis Last Modified in 01/25/19 | 208 | 159 | 157 | 0 | 5 | 0 | 0 | 2 |
[37] † Acheloma spp. Last Modified in 01/25/19 | 66 | 293 | 293 | 0 | 11 | 0 | 0 | 2 |
[38] † Eryops megalocephalus Last Modified in 01/25/19 | 54 | 305 | 304 | 0 | 12 | 0 | 0 | 2 |
[39] † Doleserpeton annectens Last Modified in 01/25/19 | 59 | 298 | 296 | 0 | 15 | 0 | 0 | 2 |
[40] Caudata Last Modified in 01/25/19 | 95 | 251 | 227 | 0 | 48 | 0 | 0 | 2 |
[41] Anura Last Modified in 01/25/19 | 112 | 216 | 206 | 0 | 52 | 0 | 0 | 2 |
[42] † Albanerpetontidae Last Modified in 01/25/19 | 214 | 131 | 130 | 0 | 26 | 0 | 0 | 2 |
[43] † Micropholis stowi Last Modified in 01/25/19 | 86 | 284 | 275 | 0 | 9 | 0 | 0 | 2 |
[44] † Eoscopus lockardi Last Modified in 01/25/19 | 90 | 277 | 272 | 0 | 8 | 0 | 0 | 2 |
[45] † Tambachia trogallas Last Modified in 01/25/19 | 167 | 197 | 196 | 0 | 7 | 0 | 0 | 2 |
[46] † Triadobatrachus massinoti Last Modified in 01/25/19 | 231 | 112 | 108 | 0 | 31 | 0 | 0 | 2 |
[47] † Gerobatrachus hottoni Last Modified in 01/25/19 | 180 | 184 | 184 | 0 | 6 | 0 | 0 | 2 |
[48] † Platyrhinops lyelli Last Modified in 01/25/19 | 101 | 261 | 260 | 0 | 9 | 0 | 0 | 2 |
[49] † Amphibamus grandiceps Last Modified in 01/25/19 | 200 | 168 | 168 | 0 | 2 | 0 | 0 | 2 |
[50] † Lethiscus stocki Last Modified in 01/25/19 | 62 | 279 | 278 | 0 | 30 | 0 | 0 | 2 |
[51] † Coloraderpeton brilli Last Modified in 01/25/19 | 76 | 261 | 259 | 0 | 35 | 0 | 0 | 2 |
[52] † Edops craigi Last Modified in 01/25/19 | 103 | 252 | 252 | 0 | 15 | 0 | 0 | 2 |
[53] † Petrolacosaurus kansensis Last Modified in 01/25/19 | 65 | 284 | 284 | 0 | 21 | 0 | 0 | 2 |
[54] † Ophiacodon uniformis Last Modified in 01/25/19 | 10 | 334 | 334 | 0 | 26 | 0 | 0 | 2 |
[55] † Baphetes spp. Last Modified in 01/25/19 | 136 | 226 | 226 | 0 | 8 | 0 | 0 | 2 |
[56] † Eucritta melanolimnetes Last Modified in 01/25/19 | 164 | 199 | 199 | 0 | 7 | 0 | 0 | 2 |
[57] † Dimetrodon spp. Last Modified in 01/25/19 | 28 | 314 | 314 | 0 | 28 | 0 | 0 | 2 |
[58] † Opisthodontosaurus carrolli Last Modified in 01/25/19 | 235 | 123 | 123 | 0 | 12 | 0 | 0 | 2 |
Taxonomic Overview for Matrix 'M25828' (61 Taxa)
taxon | unscored cells |
scored cells |
no cell support |
NPA cells |
"-" cells | cell images | labels on cell images |
member access |
[1] † Acanthostega Last Modified in 01/25/19 | 2 | 220 | 219 | 0 | 6 | 0 | 0 | 2 |
[2] † Proterogyrinus Last Modified in 01/25/19 | 20 | 207 | 203 | 0 | 4 | 0 | 0 | 2 |
[3] † Balanerpeton Last Modified in 01/25/19 | 12 | 210 | 210 | 0 | 5 | 0 | 0 | 2 |
[4] † Dendrerpeton Last Modified in 01/25/19 | 19 | 204 | 204 | 0 | 4 | 0 | 0 | 2 |
[5] † Tuditanus Last Modified in 01/25/19 | 25 | 185 | 185 | 0 | 17 | 0 | 0 | 2 |
[6] † Asaphestera Last Modified in 01/25/19 | 58 | 163 | 162 | 0 | 7 | 0 | 0 | 2 |
[7] † Hapsidopareion Last Modified in 01/25/19 | 62 | 153 | 153 | 0 | 12 | 0 | 0 | 2 |
[8] † Llistrofus pricei Last Modified in 01/25/19 | 68 | 151 | 151 | 0 | 8 | 0 | 0 | 2 |
[9] † Saxonerpeton Last Modified in 01/25/19 | 26 | 186 | 185 | 0 | 16 | 0 | 0 | 2 |
[10] † Pantylus Last Modified in 01/25/19 | 7 | 213 | 212 | 0 | 8 | 0 | 0 | 2 |
[11] † Cardiocephalus sternbergi Last Modified in 01/25/19 | 83 | 136 | 135 | 0 | 9 | 0 | 0 | 2 |
[12] † Cardiocephalus peabodyi Last Modified in 01/25/19 | 69 | 150 | 149 | 0 | 9 | 0 | 0 | 2 |
[13] † Euryodus primus Last Modified in 01/25/19 | 52 | 163 | 162 | 0 | 13 | 0 | 0 | 2 |
[14] † Euryodus dalyae Last Modified in 01/25/19 | 60 | 154 | 154 | 0 | 13 | 0 | 0 | 2 |
[15] † Pariotichus Last Modified in 01/25/19 | 140 | 80 | 80 | 0 | 7 | 0 | 0 | 2 |
[16] † Euryodus bonneri Last Modified in 01/25/19 | 136 | 84 | 84 | 0 | 7 | 0 | 0 | 2 |
[17] † UNSM 32144 Last Modified in 01/25/19 | 97 | 123 | 123 | 0 | 7 | 0 | 0 | 2 |
[18] † Tambaroter Last Modified in 01/25/19 | 112 | 107 | 107 | 0 | 8 | 0 | 0 | 2 |
[19] † Pelodosotis Last Modified in 01/25/19 | 31 | 187 | 186 | 0 | 10 | 0 | 0 | 2 |
[20] † Nannaroter Last Modified in 01/25/19 | 104 | 115 | 115 | 0 | 8 | 0 | 0 | 2 |
[21] † Micraroter Last Modified in 01/25/19 | 38 | 184 | 181 | 0 | 8 | 0 | 0 | 2 |
[22] † Rhynchonkos Last Modified in 01/25/19 | 31 | 189 | 188 | 0 | 8 | 0 | 0 | 2 |
[23] † Eocaecilia Last Modified in 01/25/19 | 33 | 168 | 167 | 0 | 27 | 0 | 0 | 2 |
[24] † Microbrachis Last Modified in 01/25/19 | 7 | 208 | 207 | 0 | 13 | 0 | 0 | 2 |
[25] † Carrolla Last Modified in 01/25/19 | 88 | 110 | 110 | 0 | 29 | 0 | 0 | 2 |
[26] † Batropetes Last Modified in 01/25/19 | 37 | 176 | 176 | 0 | 14 | 0 | 0 | 2 |
[27] † Utaherpeton Last Modified in 01/25/19 | 96 | 123 | 120 | 0 | 11 | 0 | 0 | 2 |
[28] † Stegotretus Last Modified in 01/25/19 | 49 | 172 | 170 | 0 | 8 | 0 | 0 | 2 |
[29] † Sauropleura scalaris Last Modified in 01/25/19 | 23 | 195 | 194 | 0 | 10 | 0 | 0 | 2 |
[30] † Ptyonius Last Modified in 01/25/19 | 25 | 192 | 191 | 0 | 11 | 0 | 0 | 2 |
[31] † Urocordylus Last Modified in 01/25/19 | 76 | 142 | 141 | 0 | 10 | 0 | 0 | 2 |
[32] † Keraterpeton galvani Last Modified in 01/25/19 | 39 | 179 | 179 | 0 | 9 | 0 | 0 | 2 |
[33] † Batrachiderpeton Last Modified in 01/25/19 | 43 | 180 | 177 | 0 | 7 | 0 | 0 | 2 |
[34] † Diceratosaurus Last Modified in 01/25/19 | 46 | 171 | 171 | 0 | 10 | 0 | 0 | 2 |
[35] † Diplocaulus magnicornis Last Modified in 01/25/19 | 20 | 200 | 198 | 0 | 9 | 0 | 0 | 2 |
[36] † Diploceraspis Last Modified in 01/25/19 | 47 | 170 | 170 | 0 | 10 | 0 | 0 | 2 |
[37] † Scincosaurus Last Modified in 01/25/19 | 36 | 180 | 179 | 0 | 12 | 0 | 0 | 2 |
[38] † Brachydectes Last Modified in 01/25/19 | 8 | 200 | 199 | 0 | 20 | 0 | 0 | 2 |
[39] † Adelogyrinus Last Modified in 01/25/19 | 75 | 140 | 138 | 0 | 14 | 0 | 0 | 2 |
[40] † Oestocephalus Last Modified in 01/25/19 | 18 | 151 | 151 | 0 | 58 | 0 | 0 | 2 |
[41] † Phlegethontia Last Modified in 01/25/19 | 22 | 123 | 122 | 0 | 83 | 0 | 0 | 2 |
[42] † Greererpeton Last Modified in 01/25/19 | 10 | 214 | 210 | 0 | 7 | 0 | 0 | 2 |
[43] † Seymouria baylorensis Last Modified in 01/25/19 | 9 | 217 | 214 | 0 | 4 | 0 | 0 | 2 |
[44] † Limnoscelis Last Modified in 01/25/19 | 13 | 210 | 209 | 0 | 5 | 0 | 0 | 2 |
[45] † Branchiosauridae Last Modified in 01/25/19 | 50 | 172 | 169 | 0 | 8 | 0 | 0 | 2 |
[46] † Micromelerpetontidae Last Modified in 01/25/19 | 54 | 172 | 166 | 0 | 7 | 0 | 0 | 2 |
[47] † Tersomius Last Modified in 01/25/19 | 84 | 147 | 140 | 0 | 3 | 0 | 0 | 2 |
[48] † Ecolsonia Last Modified in 01/25/19 | 49 | 176 | 174 | 0 | 4 | 0 | 0 | 2 |
[49] † Acheloma Last Modified in 01/25/19 | 80 | 143 | 143 | 0 | 4 | 0 | 0 | 2 |
[50] † Eryops Last Modified in 01/25/19 | 67 | 155 | 155 | 0 | 5 | 0 | 0 | 2 |
[51] † Doleserpeton Last Modified in 01/25/19 | 57 | 170 | 167 | 0 | 3 | 0 | 0 | 2 |
[52] Salamanders Last Modified in 01/25/19 | 7 | 191 | 177 | 0 | 43 | 0 | 0 | 2 |
[53] Frogs Last Modified in 01/25/19 | 9 | 175 | 168 | 0 | 50 | 0 | 0 | 2 |
[54] † Albanerpetontidae Last Modified in 01/25/19 | 81 | 119 | 118 | 0 | 28 | 0 | 0 | 2 |
[55] † Micropholis Last Modified in 01/25/19 | 23 | 206 | 199 | 0 | 5 | 0 | 0 | 2 |
[56] † Eoscopus Last Modified in 01/25/19 | 26 | 200 | 195 | 0 | 6 | 0 | 0 | 2 |
[57] † Tambachia Last Modified in 01/25/19 | 79 | 145 | 144 | 0 | 4 | 0 | 0 | 2 |
[58] † Triadobatrachus Last Modified in 01/25/19 | 90 | 108 | 104 | 0 | 33 | 0 | 0 | 2 |
[59] † Gerobatrachus Last Modified in 01/25/19 | 89 | 134 | 134 | 0 | 4 | 0 | 0 | 2 |
[60] † Platyrhinops Last Modified in 01/25/19 | 54 | 168 | 168 | 0 | 5 | 0 | 0 | 2 |
[61] † Amphibamus Last Modified in 01/25/19 | 48 | 176 | 176 | 0 | 3 | 0 | 0 | 2 |
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Matrix downloads | 4 | Pardo et al 2017 matrix with recoded Llistrofus (2 downloads); Huttenlocker et al 2013 matrix with recoded Llistrofus (2 downloads); |